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RESEARCH COMMUNICATIONS
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ACKNOWLEDGEMENTS. We thank DST, New Delhi for financial
support (Grant No: SR/S4/ES-389/2008). We also thank the anonymous
reviewers for their incisive comments that helped improve the manuscript.
Interactions of lion-tailed macaque
(Macaca silenus) with non-primates in
the Western Ghats, India
Joseph J. Erinjery 1, Honnavalli N. Kumara2 ,
K. Mohan1 and Mewa Singh1,3,4,*
1
Bio-Psychology Laboratory and Institute of Excellence,
University of Mysore, Mysuru 570 006, India
2
Sálim Ali Centre for Ornithology and Natural History,
Coimbatore 641 108, India
3
Evolutionary and Organismal Biology Unit, Jawaharlal Nehru
Centre for Advanced Scientific Research, Bengaluru 560 064, India
4
National Institute of Advanced Studies, Indian Institute of
Science Campus, Bengaluru 560 012, India
Primates and non-primates inhabiting tropical forests
may interact with each other since they coexist in the
same communities. Primates usually interact with
their prey, predators, competitors and neutral species.
Using ‘all occurrence’ sampling, we have studied
inter-specific interactions of lion-tailed macaques with
non-primate species found in their habitat. We observed that the percentage of total time spent on interactions with non-primates was less than 1. Also, the
percentage of total time spent in interacting with
competitors, predators and neutral species was less
than 0.5. The lack of predation pressure and lack of
opportunities for mixed-species associations for increasing foraging efficiency appear to be the major
reasons for the absence of interactions with nonprimates. By comparing with studies from other primate habitat regions, we observed that primates in
South Asia interact much lesser with non-primates
than those in South America and Africa. A previous
study showed that the interactions of lion-tailed macaques even with other primate species in the Western
Ghats are less than expected by chance.
Keywords: Inter-specific competition, mixed-species
troops, Macaca silenus, primate–predator interaction.
PRIMATES in tropical forests may interact with other nonprimate species which coexist with them1–3. These interactions could be either positive, negative or neutral4,5, and
may vary with space and time5–7, largely depending on
ecological and historical factors which shape them. The
primates mainly interact with their prey, predators, feeding competitors and neutral species, and may display a
variety of behaviours towards an interacting species5.
Most direct interactions such as aggression and submission occur when two species encounter in close proximity8. Long-distance interactions such as alarm calls and
spacing calls are important for communication about
detection of predators and foraging9.
Received 30 July 2016; revised accepted 29 November 2016
doi: 10.18520/cs/v112/i10/2122-2129
*For correspondence. (e-mail: [email protected])
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The primate–non-primate interactions are more common in the neotropics, followed by Africa and Asia 3. The
primate–non-primate interactions are mainly with other
mammals and birds, and usually the non-primates are attracted to the primates than the other way around3. It is
stipulated that the chances of such commensal interactions are high in Asia, Africa and the neotropics, as primates in these regions are mainly large/medium-bodied,
wasteful feeders with large group sizes inhabiting open
habitats3. However, it has been observed that both body
size and group size do not have any effect on primate–
non-primate interactions3. The nature of primate interactions could be largely determined by species-specific
temperament and personality5,10,11.
Some interactions after encounters may result in active
associations between interacting species which are
formed more commonly for reducing predation pressure
and increasing foraging efficiency1,6,12. Generally, two
species are defined to be in an association when they are
in close proximity, usually within a distance of 50 m, and
this may vary with the species being studied and their
habitats13. Information flow between species that share
common resources and predators is an important aspect
of communities9. Such associations have been largely observed among primate–non-primate species as in the case
of Hanuman langur (Semnopithecus entellus) – chital
(Axis axis)14 and chacma baboon (Papio ursinus) – rock
kestrel (Falco rupicolus) associations7. Also, the animals
are often observed to interact when feeding on clumped
resources and when their travel routes converge15.
Lion-tailed macaques are habitat specialists of tropical
rainforests of the Western Ghats, India16–18. Previous
studies have shown that lion-tailed macaques avoid interacting with other primate species in the Western Ghats11,19.
Also, the studies assessing the inter-specific interactions
of primates with non-primates is South Asia are rare in
the literature. However, such studies are essential to understand community ecology and interdependencies of
species in an ecosystem. Hence, we studied inter-specific
interactions of lion-tailed macaques with other nonprimates to understand whether primates in the Western
Ghats interact with other species. Based on previous studies conducted on inter-specific interactions of other primates in Asia and primate–primate interactions in the
Western Ghats11,13,20 , we expected the lion-tailed macaque
to spend less time on inter-specific interactions with other
non-primates in the Western Ghats. As lion-tailed macaques are found both in highly fragmented forests with
largely open canopies and human disturbance as well as
relatively undisturbed and large stretches of continuous
forests21, we also intended to compare such inter-specific
interactions between groups inhabiting both habitat types.
Our study conformed to the ethical guidelines for animal
treatment of the University of Mysore, and we obtained
permissions for conducting wildlife research from the
Forest Departments of Kerala and Tamil Nadu.
2130
We conducted the study in two sites in the Western
Ghats: Nelliyampathy and Valparai. In Valparai, data
were collected from two different locations: Andiparai
and Puthuthottam. Details about the study groups in Nelliyampathy and Andiparai are given in Erinjery et al.11.
The group in Nelliyampathy inhabited a contiguous forest
surrounded by coffee/tea gardens and the groups in Andiparai and Puthuthottam inhabited fragmented forest
patches. Puthuthottam is a privately owned rainforest
fragment of about 1.25 km2 area surrounded by tea gardens, at an altitude of about 1100 m. Since both these are
fragmented habitats, we combined the data of these locations for the present study.
The study in Nelliyampathy was conducted from December 2010 to October 2012 with 736 h of total observation time. The study in Valparai was conducted from
September 1996 to August 1997 with 1696 h of total observation time. We systematically followed the macaques
for 92 and 212 d in Nelliampathy and Valparai respectively (8 h/day), and used only these data for the analysis.
The data were collected between 0800 and 1600 h for full
day.
We employed ‘all occurrences’ sampling method by
taking detailed notes of interactions, as and when they
occurred. We collected data on date, time, interacting
species, activity of macaque and other species during the
interaction, type of interaction, aggressor and the recipient, displacement (if any), species displaced, and duration
of the interaction. For analysis, we considered only those
interactions wherein we could observe the beginning and
end of an interaction.
We divided inter-specific interactions into two types:
long-distance interactions and close-proximity interactions. We included alarm calls as a long-distance interaction. We considered an arboreal species to be involved in
a close-proximity interaction only if that species was
within a proximity of 5 m from an individual of the liontailed macaque group. For terrestrial animals, we considered an interaction to take place only when the animals
were directly under the trees where lion-tailed macaques
were present. We considered the lion-tailed macaque to
interact with a predator, which was mostly a raptor, only
when the latter directly hovered over the trees where liontailed macaques were present. We further divided the
close-proximity interactions into two types: neutral and
non-neutral. In neutral interactions, lion-tailed macaques
tolerated the other species. In non-neutral interactions,
they were aggressive towards other species. Aggressive
interactions included chasing, fighting and/or displacing
of interacting species. When interacting with predators,
we considered lion-tailed macaques to be aggressive if
they gave continuous threatening bark calls directed at
the predator. We did not observe any interactions where
lion-tailed macaques were submissive to other species.
On the basis of previous studies on lion-tailed
macaques and our own long-term observations in the
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Table 1.
Frequency, mean duration, percentage of total frequency and percentage of total time spent on inter-specific interactions by lion-tailed
macaques with non-primates in Nelliyamapathy and Valparai
Species
interacting
Location
Frequency
of interaction
Mean duration of an
interaction (min)
Percentage of total
frequency of interaction
Percentage of total time
spent on interaction
Nelliyampathy
DHL*
SE, MHE*
GS*
GIH*
RTD*
SBR*
Total
2
6
24
6
14
1
53
7.5
11.6
6.3
5.0
9.6
5.0
7.6
3.8
11.3
45.3
11.3
26.4
1.9
100
3.7
17.3
37
7.4
33.3
1.2
100
Valparai
DHL
SE
LEO*
GS
GIH
BD*
RTD
SBR
ELE*
Total
2
18
1
34
3
7
8
3
1
77
5.0
11.7
30.0
17.9
16.7
17.9
9.8
5.0
15
12.51
2.6
23.4
1.3
44.2
3.9
9.1
10.4
3.9
1.3
100
1.0
21.8
3.1
44.6
5.2
13.0
8.1
1.6
1.6
100
*GS, Indian giant squirrel; DHL, Dhole; SE, Crested-serpent eagle; MHE, Mountain hawk-eagle (only one killing episode); RTD, Racket tailed
drongo; SBR, Sambar; GIH, Great Indian hornbill; BD, Barking deer; LEO, Leopard; ELE, Elephant.
Figure 1.
as prey.
A mountain hawk-eagle with a lion-tailed macaque infant
field, we categorized the results of interactions as follows: interactions with predators, interactions with competitors and interactions with commensal/mutualistic
species. The major predators of lion-tailed macaques
were leopard (Panthera pardus), crested serpent eagle
(Spilornis cheela) and changeable hawk-eagle (Nisaetus
cirrhatus)22,23. However, we also included other species
which elicited threat calls and alarm calls from lion-tailed
macaques. The species which shared similar food resources with lion-tailed macaques were considered as
CURRENT SCIENCE, VOL. 112, NO. 10, 25 MAY 2017
competitors. We considered Indian giant squirrel (Ratufa
indica) and great Indian hornbill (Buceros bicornis) as
competitors, since they shared the same fruits that the
macaques consumed. If the other species benefitted from
the interaction with lion-tailed macaques, we considered
it as commensal and if both species benefited, we considered it a mutualistic interaction.
We used chi-square test of proportions to compare between interactions from different study sites. The alpha
level was kept at 0.05 and we performed all tests using
SPSS 20.
Lion-tailed macaques spent 0.92% and 0.95% of their
time in interacting with non-primates in Nelliyampathy
and Valparai respectively, with no difference in the duration of interactions across study sites (chi-square test of
proportions:  2 = 0.051, P = 0.48). The frequency of
interactions was 0.05 interactions/h in Valparai and 0.07
interactions/h in Nelliyampathy (Table 1).
The interactions with non-primate species included
0.19% and 0.26% of their time with potential predators
in Nelliyampathy and Anamalai respectively, with no
difference across sites (chi-square test of proportions:
 2 = 0.051, P = 0.17). Out of the total time spent on
interactions, lion-tailed macaques interacted with their
predators 21% and 27.5% of their time in Nelliyampathy
and Valparai respectively (Table 1). We observed two
killing episodes by the predator (a lion-tailed macaque
infant caught by crested serpent eagle and the other by
mountain hawk-eagle, Nisaetus nipalensis) during the
whole study period. The observation of mountain hawkeagle capturing lion-tailed macaque infant is the first case
of such predation in the wild (Figure 1). The crested
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Table 2.
Frequency and mean time spent on neutral and non-neutral interactions by lion-tailed macaques (LTMs) with non-primates in
Nelliyampathy and Valparai. Values in parenthesis are the mean time spent on each interaction
Nelliyampathy
Valparai
Type of interaction
Frequency (mean time spent (min)/
interaction)
Interaction
LTM and DHL
LTM and SE, MHE
LTM and LEO
LTM and GS
LTM and GIH
LTM and RTD
LTM and SBR
LTM and BD
LTM and ELE
Observation
time (h)
Aggressive
Tolerant
736
736
736
736
736
736
736
736
736
2 (7.5)
6 (11.6)
0 (0)
11 (8.63)
1 (5)
4 (5)
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
13 (9.6)
5 (6)
10 (11.5)
1 (5)
0 (0)
0 (0)
Submissive
0
0
0
0
0
0
0
0
0
serpent eagle was a common predator for primates. They
continuously circled around and swooped down when
presented with an opportunity. The lion-tailed macaques
gave repeated alarm calls at the sight of crested serpent
eagle and mountain hawk-eagle (N = 24). Twenty out of
23 interactions with crested serpent eagles took place during the months (June–August) when the macaques had
young infants. There were four instances of interactions
with dholes (Cuon alpinus), and lion-tailed macaques
made barking/threatening calls at the sight of dholes. The
lion-tailed macaques also made barking calls in the presence of Asian elephants (Elephas maximus; N = 1) and
leopard (N = 1).
The most frequent non-primate interaction of liontailed macaques was with the Indian giant squirrel
(N = 58). The macaques spent 0.43% (0.03 interactions/h)
and 0.41% (0.02 interactions/h) time interacting with
squirrels in Nelliyampathy and Valparai respectively. Out
of the total time spent on interactions, lion-tailed macaques spent 44.6% and 37% of their time interacting with
Indian giant squirrel alone in Valparai and Nelliyampathy
respectively. Table 2 presents the data on the type of interactions of lion-tailed macaques with the other species.
The lion-tailed macaques were more frequently neutral
(N = 43) than aggressive towards Indian giant squirrel
(N = 15). Most aggressive interactions (93%) occurred
when the two species were feeding. Lion-tailed macaques
and other primates in the study area, Nilgiri langurs
(Semnopithecus johnii) and bonnet macaques (Macaca
radiata) became alert whenever Indian giant squirrels
gave alarm calls. Only two out of nine interactions with
the great Indian hornbill included aggression. All these
interactions occurred on Ficus trees, where both were
seen feeding together on figs.
Macaques spent 0.30% (0.02 interactions/h) and 0.17%
(0.01 interactions/h) of their time interacting with rackettailed drongos (Dicrurus paradiseus) in Nelliyampathy
2132
(0)
(0)
(0)
(0)
(0)
(0)
(0)
(0)
(0)
Type of interaction
Frequency {mean time spent (min)/
interaction}
Observation
time (h)
Aggressive
Tolerant
1696
1696
1696
1696
1696
1696
1696
1696
1696
2 (5)
18 (11.7)
1 (30)
4 (7.5)
1 (15)
2 (5)
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
0 (0)
30 (13.3)
2 (15)
6 (11.4)
3 (5)
7 (17.9)
1 (15)
Submissive
0
0
0
0
0
0
0
0
0
(0)
(0)
(0)
(0)
(0)
(0)
(0)
(0)
(0)
and Valparai respectively. These interactions took place
mainly when the macaques were foraging/feeding for insects (N = 22). The drongos continuously followed the
macaques and fed along with them. In 6 out of 22 interactions, the macaques chased the drongos. All interactions
were commensal. Macaques spent 0.01% (<0.01 interactions/h) and 0.03% (<0.01 interactions/h) time interacting
with sambar (Rusa unicolor) in Nelliyampathy and Valparai respectively. Both were found to feed together on
jackfruit (Artocarpus heterophyllus) during monsoon
(N = 4). The macaques reacted with alertness to the alarm
calls given by the sambar in three observations. Barking
deer (Muntiacus muntjak) were observed to feed on jackfruit dropped by lion-tailed macaques (N = 5), and the
macaques reacted with alertness to the alarm calls by
barking deer (N = 4). The interactions with barking deer
were only seen in Valparai (0.28%; 0.01 interactions/h).
The interdependencies of species in a community are
important to understand the community ecology of a species. While several researchers have studied polyspecific
associations and interactions between primate species,
studies describing the non-primate interactions in South
Asia are rare in the primate literature. In the present
study, we collected data on non-primate inter-specific
interactions of an endangered primate, the lion-tailed
macaque, inhabiting the Western Ghats of South India, to
assess their interdependency with non-primates. We
found that the interaction of lion-tailed macaques with
non-primates was <1% of the total observation time. This
shows that the lion-tailed macaque hardly interacts with
any non-primate in its habitat. These results are similar to
the primate–primate interactions in the Western Ghats11.
Haugaasen and Peres1 showed that interactions
between primates and other mammals are rare in central
Amazonian forests. However, a meta-analysis from different regions has shown that primate–non-primate interactions are most common in the neotropics, followed by
CURRENT SCIENCE, VOL. 112, NO. 10, 25 MAY 2017
RESEARCH COMMUNICATIONS
Africa and Asia3. It has been observed that the interactions between primates and other species in Africa,
Southeast Asia and Central America are higher compared
to the present study1,7,24–28. The absence of large raptors
that predate on adult primates may be the major reason
for absence of associations or interactions between primates and non-primates in South Asia29.
Primates mainly form associations after encounters
with other non-primates for increasing foraging efficiency or decreasing predation pressure3,29. The present
study has shown that the predation pressure for liontailed macaques is less in the Western Ghats (<0.50% of
the total observation time), and they do not associate with
other non-primates for increasing foraging efficiency. It
has been observed that the lion-tailed macaques are
aggressive towards Indian giant squirrel in most of the
interactions during co-feeding. This is probably because
of competition between these species, which feed on
fruits of the same trees. The lion-tailed macaque has even
been observed to predate on the Indian giant squirrel,
Indian giant flying squirrel (Petaurista philippensis),
three-striped squirrel (Funambulus palmarum), frogs, lizards and bats16,30–32. One instance of lion-tailed macaque
unsuccessfully attacking a mouse deer (Moschiola indica)
fawn is also reported30. However, it has been found that
other mammals and birds, including racket-tailed drongo,
barking deer and sambar were associated with lion-tailed
macaques to increase their foraging efficiency by feeding
on the food dropped by the macaques. Also, lion-tailed
macaques were observed to be alert to alarm calls of
barking deer, sambar and Indian giant squirrel. Hence,
while interactions with racket-tailed drongos were
commensal, those with barking deer and sambar were
mutualistic. Additionally, we observed the Puthuthottam
lion-tailed macaque group to interact with humans frequently.
The lack of inter-specific interactions in different habitats of lion-tailed macaques indicates that the community
composition of higher vertebrates in the Western Ghats
might not be largely influenced by interdependencies between species, but due to the anthropogenic and historical
factors in the past. Also, the low percentage of time observed for associating and interacting shows that liontailed macaques inhabiting the Western Ghats do not
form active associations with other vertebrates, with only
chance interactions due to sharing of similar habitats with
other species. However, the frequency of interactions
may depend on the density and abundance of nonprimates in a particular study site. Also, the present study
shows that there is little difference in interactions between lion-tailed macaques inhabiting contiguous and
fragmented forests.
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Received 4 July 2016; revised accepted 12 December 2016
India and China participate in the global market. This
communication focuses on intelligent interacting systems which are present over globe and these are a
source of rising the software development cycle with
the help of modern communication facilities. Free
e-Market globalization is now vital for billions of people. However, IT leadership is not possible without a
review of the existing system. The present study is
based over the issue of the global market related research, education and investment in IT technology.
IT-based-leadership can give sustainable global competitive advantage to our country. So the role of iterative software development is crucial to be targeted in a
systematic fashion.
Keywords: Global software development, information
technology, innovation, software services, technology.
T HE developing countries are adopting incremental
export-oriented software development. These services
and trends of the industry are the basis for talent and
quality of work, and have also brought more investment.
Europe and America are high-wage countries for software
and services; they are now increasingly looking for cheap
labour, thus resulting in offshoring. International trading
of software services creates several jobs. Services trade in
software development provides more skilled workers to
firms of developing countries. This aspect leads the policy makers to target more opportunities and challenges to
produce precious national income. This income and new
employment opportunities are the real fruit of the global
software development (GSD)1–3. Multinational companies
are doing business in countries with low wages for capital
saving though local markets.
In the present study, we consider the following:
doi: 10.18520/cs/v112/i10/2129-2134
Interactive systems regarding global
software development and offshoring
Ramzan Talib1 , Muhammad Yahya Saeed1,*,
Muhammad Awais2 and Kashif Hanif1
1
Department of Computer Science, and
Department of Software Engineering, Govt College University
Faisalabad, Illama Iqbal Road, Faisalabad 38000, Pakistan
2
Modern information technology (IT) methods are reshaping the global market with great success. With
today’s global software industry, IT has made innovations everywhere, including businesses and consumer
practices. This has made developing countries like
*For correspondence. (e-mail: [email protected])
2134
(a) Software and information technology (IT) as a
whole.
(b) Programming and development.
(c) Software testing of all types using the principles of
GSD.
(d) Remotely performing software maintenance work
that is offshore.
(e) On-line research and development, such as software
architecture, product design, project management,
etc.
(f) IT consultation and on-line guidance of business
strategy.
(g) Physical product manufacturing like semiconductors,
computer hardware, etc.
(h) Business process outsourcing/services such as financial analysis, on-line accounting services, digital art,
desktop publishing, high-end services, etc.
(i) Call centres and telemarketing.
The software industry works across national borders
more closely associated with all of the above categories.
CURRENT SCIENCE, VOL. 112, NO. 10, 25 MAY 2017